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sustainability Article Contrasting Effects of Tillage and Landscape Structure on Spiders and Springtails in Vineyards Alexandra Pfingstmann 1, Daniel Paredes 1,2 , Jacob Buchholz 1, Pascal Querner 1, Thomas Bauer 3, Peter Strauss 3 , Sophie Kratschmer 4 , Silvia Winter 4,5 and Johann Zaller 1,* 1 Institute of Zoology, University of Natural Resources and Life Sciences Vienna (BOKU), A-1180 Vienna, Austria; alexandra.pfi[email protected] (A.P.); [email protected] (D.P.); [email protected] (J.B.); [email protected] (P.Q.) 2 Grupo de Protección Vegetal, Departamento de Protección Ambiental, Estación Experimental de Zaidín, CSIC, 18008 Granada, Spain 3 Institute for Land and Water Management Research, Austrian Federal Agency for Water Management, A-3252 Petzenkirchen, Austria; [email protected] (T.B.); [email protected] (P.S.) 4 Institute of Integrative Nature Conservation Research, University of Natural Resources and Life Sciences Vienna (BOKU), A-1180 Vienna, Austria; [email protected] (S.K.); [email protected] (S.W.) 5 Division of Plant Protection, University of Natural Resources and Life Sciences Vienna (BOKU), A-1180 Vienna, Austria * Correspondence: [email protected]; Tel.: +43-1476-5483-318 Received: 4 February 2019; Accepted: 30 March 2019; Published: 8 April 2019 Abstract: Interactions between predatory species and their potential prey are little studied in vineyards, especially considering the surrounding landscape structure. We examined the effects of soil tillage intensities in vineyard inter-rows on the activity density and diversity of spiders (Araneae) and springtails (Collembola), their potential preys, and assessed whether these effects are altered by non-crop elements in the surrounding landscape. We collected data in 16 vineyards in Austria; eight were periodically mechanically disturbed (PMD), eight had permanent green cover (PGC). The study vineyards were embedded in landscapes ranging from structurally simple to complex. Both, spiders and springtails were collected with pitfall traps. Data analyses using generalized linear mixed models (GLMM) showed different effects of soil tillage intensities on spiders and springtails and an interaction with semi-natural elements (SNEs) in the surrounding landscape. Activities of springtails were higher under PMD than under PGC while spider activity density remained unaffected. Spider family Shannon diversity was lower under PMD than under PGC, while springtail species Shannon diversity was unaffected by tillage. Under PMD, spider activity and family diversity decreased with increasing SNEs in the surroundings indicating spider emigration away from vineyards. Under PGC, spider activity density increased with increasing SNE proportions in the surroundings when springtail activity density was high. Our findings suggest that recommendations on sustainable vineyard management should include both site and landscape factors. Keywords: agroecology; viticulture; soil cultivation; landscape ecology 1. Introduction Viticulture is a very old sector of agriculture, dating back to about 6000 BC [1]. Modern conventional vineyards are usually intensively managed with frequent use of pesticides and tillage during the vegetation period [2]. Inter-row soil management practices are imposed to control weeds Sustainability 2019, 11, 2095; doi:10.3390/su11072095 www.mdpi.com/journal/sustainability Sustainability 2019, 11, 2095 2 of 14 and can include intensive tillage (bare soil), alternate tillage where every second inter-row is tilled, or no tillage with permanent green cover, or by the use of herbicides to control weeds [3]. Inter-row soil management in vineyards has been shown to affect aboveground [4] and belowground [5,6] biodiversity and associated ecosystem services [7]. However very little is known about influences on organisms comprising different trophic levels that might affect predator-prey interactions. Spiders (Araneae) are all predators comprising the largest order of Arachnids with around 46,800 species worldwide [8]. Their habitat requirements are defined by several abiotic (temperature, humidity, wind, light intensity) and biotic (type of vegetation, food supply, intra-and interspecific competition, predators) factors [9,10]. Spiders live in different vegetation layers and are often divided in two functional groups: Sedentary web spiders and non-web builders. Spider abundances can vary considerably in different ecosystems with mean abundances in temperate climate in grassland 2 2 2 with 92 ind. m− , forests with 53 ind. m− , and arable fields of 21 ind. m− [11]. Spiders are important generalist predators in many ecosystems [9,12,13], however their abundance and diversity is commonly decreased in managed agroecosystems [10,14,15]. While spider communities, their feeding behaviors and possible implications for biological control have frequently been investigated in arable agroecosystems [16,17], far less is known from vineyards [13,18–21]. Springtails (Collembola) are among the most widespread terrestrial microarthropods with about 8800 species worldwide representing an important component of soil fauna in many terrestrial ecosystems [22]. Springtail species diversity is usually affected by land use [15,23]. Springtails can be grouped into soil dwelling (eu- and hemiedaphics) and surface dwelling species (epedaphics) [24]. As members of the decomposer community they are feeding on fungi, bacteria and dead organic matter thereby mobilizing nutrients [25]. Abundance of springtails can vary considerably between different 2 ecosystems [11,26], ranging from abundances in grassland of 29,545 ind. m− , forests of 1826 ind. 2 2 m− to arable fields of 6056 ind. m− [11]. Additionally, land management intensity including tillage, residue management or fertilization [27,28], pesticide use [29]. Springtails are also an important source of prey to spiders and comprise up to 72% of their diets [16,30–32]. Furthermore, springtails provide essential nutrients and amino acids in the diet of spiders [33,34]. Few studies have investigated the effect of management practices in vineyards on spiders [18,20,35,36] or springtails [5]. Tillage of vineyard inter-rows commonly has negative effects on the abundance of soil invertebrates, although rapid recovery of specific invertebrate groups (e.g., earthworms) has been shown [6]. Studies conducted in arable and viticultural landscapes that assessed combined effects of management and the surrounding landscape on spiders [37–40] or springtails [5] found that complex landscapes contain a higher biodiversity than simpler structured landscapes [41]. However, spiders appeared to respond differently to landscape structure: Ambush and habitat specific spiders are more abundant in diverse landscapes whereas web builders are more abundant in simple landscapes [38]. Species richness of spiders was significantly enhanced in wheat fields with more non-crop habitats in the surroundings [17,42,43], although opposite responses have also been reported [44]. Also, springtail populations showed a response to land-use intensity and the surrounding landscape structures [5,45,46]. However, to the best of our knowledge no study investigated both spiders and springtails in vineyards in response to the surrounding landscape. The aims of the current study were (i) to assess the response of spider and springtail activity and diversity in vineyards to periodically mechanically-disturbed (PMD) versus permanently green covered (PGC) inter-rows, (ii) to analyze whether the proportion of semi-natural habitats in the surrounding landscape and the presence of springtails alter responses of spiders and (iii) to assess spider family assemblages and their relation with local management and potential springtail prey. We hypothesized that permanent green cover will benefit both spiders and springtails because of less disturbance. A higher proportion of non-crop habitats in the surroundings is expected to increase the diversity and activity of spiders and springtails in vineyards due to migration into vineyards. Sustainability 2019, 11, 2095 3 of 14 2. Materials and Methods 2.1. Site Description and Study Design The study sites were located in the two neighboring wine regions Carnuntum (48◦ 40 N, 16◦ 470 E) and Neusiedlersee-Hügelland (47◦ 540 N, 16◦ 410 E) about 40 km south east of Vienna, Austria. The annual total precipitation for the region Bruckneudorf (which is located in the center of the study regions) is around 619 mm at an altitude of 167 m and an annual average temperature of 10.2 ◦C[47]. The study regions are strongly shaped by a Pannonian, continental-like climate with hot, dry summers and cold winters. The most frequent soil types are Chernozems and Cambisols with medium to high usable field capacity, neutral pH-values and high nutrient contents [48]. Within the regions, 16 vineyards were selected for the current investigations (Figure1). In these regions vineyards are small (0.4–1 ha) and typically surrounded by landscapes with various degrees of heterogeneity. Landscape heterogeneity was calculated as the proportion of semi-natural habitats (hedges, tree rows, grass stripes, natural grassland, pasture, fallow, heathland, wetland and woodlots) within a radius of 750 m centered in the plot. This area is within the activity ranges of both spiders [17,49] and springtails [46] in agricultural landscapes. In eight of the study vineyards, the inter-rows had a permanent green cover (PGC) whereas in eight vineyards every second inter-row
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